Early development of the cephalic skeleton in the turbot

At hatching Scophthalmus maximus shows no cartilaginous and no bony structure. Mecke?s cartilages appear when the fry are 1 day old, followed on day 2, by formation of the trabecular bars, fused at the outset to form a trabecula communis. Concurrently, the palatoquadrates complete the mandibular arch, and the first two pairs of ceratobranchials, associated with a pair of hyoid bars, form the beginnings of the hyobranchial system. By day 3, the parachordals have fused with the trabecular bars, the hyosymplectics have linked to the hyoid bars by interhyals, and the first four pairs of ceratobranchials have appeared. The first bony structures appear: the preoperculars. On day 8, the frontals develop above the orbits and the maxillaries and dentaries appear. On day 10, the primordia of the taeniae marginales appear, the palatoquadrates bear a pterygoid process, and to the branchial basket have been added the fifth pair of ceratobranchials and the four pairs of epibranchials. On day 12, both pairs of posterior pharyngobranchials are present. The premaxillaries develop in front of the maxillaires, and retroarticulars and the angulars complete the lower jaws. On day 13, a thin parasphenoid contributes to the floor of the neurocranium, and ectopterygoids and entopterygoids to the splanchnocranium. The set of opercular bones is complete. On day 15, the tectum synoticum closes the braincase posteriorly. The splanchnocranium possesses a basihyal and the pharyngobranchials of the first epibranchials. On day 18, the tectum posterius completes the dome of the braincase. The rear end and lateral walls of the skull are formed by the basioccipital, the exoccipitals, the pterotics, and the parietals. The suspensorium is nearly complete. From day 10, the first resorptions begin in parallel with the construction of the chondrocranium. Mecke?s cartilages each split in two, then the posterior part of the trabecular bars disappears. On day 23, the right taenia marginalis separates from the lamina orbitonasalis and curves towards the centre. Simultaneously, the right eye begins its migration to the left. This is the only metamorphosis-linked asymmetry to appear during the development of the chondrocranium. On day 25, many more bony structures appear, a characteristic of this stage: the nasals, lateral ethmoids, mesethmoid, sphenotics, prootics, pleurosphenoids, epiotics, and supraoccipital. From this stage on, the bony structures continue to develop, while the front of the neurocranium and the jaws undergo a deep remodelling due to metamorphosis. The left taenia marginalis does not appear reduced until day 29. By day 45, there remain only a few small elements of the cartilaginous skull.     

鲂鱼的头骨发育及其适应意义

本文对鲂鱼（Ｍｅｇａｌｏｂｒａｍａｓｋｏｌｋｏｖｉｉ）头骨的早期发育过程及其与鱼苗的存活功能需要之间的关系进行了研究。头骨发育的全过程可划分为５个阶段,即软颅阶段、咽颅膜骨附加阶段、脑颅开始骨化阶段、脑颅快速骨化阶段和骨化完成阶段。刚出膜仔鱼头部即有软骨存在,最先出现的硬骨是膜质上颌骨和主鳃盖骨,脑颅最先开始骨化的是基枕骨和侧枕骨,随后才是副蝶骨。头骨发育过程与鱼苗早期存活的功能需要之间有着密切的关系。     

Development of the bony skull in common sole: brief survey of morpho-functional aspects of ossification sequence

The postembryonic development of the bony cephalic skeleton in the common sole Solea solea , observed from hatching to the juvenile stage or postmetamorphic larva, appears to follow a similar chronological order to that observed in other Pleuronectiformes and Perciformes and the sequence in bone formation is a response to functional demands. At hatching, S. solea has no bony structure. On day 4, only the outlines of maxillaries and opercular bones are visible. On day 6, a thin parasphenoid appears between the orbits and isolates the braincase from the buccal cavity making food ingestion possible without any impact on the brain. On day 8, the dentaries form and two small preopercular bones appear on each side of the head. On day 9, at weaning from the yolk sac, branchial arches support the gill filaments (used for respiration and trapping phytoplankton which pass through the open mouth). On day 10, the premaxillaries develop in front of the maxillaries. The superimposing of the maxillaries and the premaxillaries is a typical feature of species possessing an acanthopterygian protractile mouth at the adult stage. On day 12, the frontals develop above the orbits and the set of opercular bones is complete. On day 18, the migration of the left eye begins. On day 20, the left eye has moved to the median crest of the head. On day 23, both eyes are located on the same side. On day 26, the braincase is formed by a basioccipital, exoccipitals, pterotics, sphenotics and a supraoccipital. On day 50, new structures have appeared, others have developed and have undergone an extensive remodeling due to metamorphosis.     

Osteocranial development in the viviparous surfperch Amphistichus argenteus (pisces: Embiotocidae)

The development of the cranial and branchial skeleton of the surfperch Amphistichus argenteus, a member of the family Embiotocidae, is described, and phylogenetic and functional aspects of the skull development of this species are discussed. The earliest bones to appear are those dermal elements of the branchial skeleton involved with feeding, and the bones, both dermal and endochondral, located in the basicranial region of the neurocranium. These are followed by dermal bones associated with the lateral line system and finally by the remainder of the bones of the branchial skeleton and the cartilaginous bones of the otic capsules. The last bone to develop is the ethmoid.     

Early development of the skull of Sander lucioperca (L.) (Teleostei: Percidae) relating to growth and mortality

The early osteological development of the skull (chondrocranium and osteocranium) of the pikeperch Sander lucioperca was studied. Specimens were reared at two temperatures, 15·5 and 18·0° C, from hatching until 47 and 43 days after fertilization (DAF), respectively. The skeletal elements characteristic for the different developmental stages were the same at both rearing temperatures, but pikeperch reared at 15·5° C reached the developmental stages later. The formation of the functional complexes, the neurocranium, jaws and suspensorium, branchial basket and hyoid arch, was evaluated chronologically. The focus was on skull development during several functional changes: at hatching, at the shift from endogenous to mixed feeding, the shift to exclusively exogenous feeding and upon reaching the final prey-capture mechanism. Growth in total length differed between fishes reared at the two temperatures, except during a phase of very slow growth from the end of the embryonic stage until the second larval stage. The latter phase, in which most of the bony elements of the viscerocranium started to form, was marked by high mortality. When exogenous feeding began, the growth rates at both temperatures increased distinctly and the first bony elements were formed in the neurocranium. Specimens reared at 18·0° C grew continuously, but those at 15·5° C showed a second period of slow growth and high mortality. Fish reared at 18·0° C reached the successive larval stages distinctly earlier than fish reared at 15·5° C.     

Early development of the chondrocranium in Chrysichthys auratus

The inception and development of the cartilaginous cephalis skeleton of Chrysichthys auratus is described from hatching to about 18 days post-hatching. At hatching, no skeletal structure is present. Not until day 3 do clearly delimited cranial primordia become apparent. As in many siluriforms, the neurocranium is platybasic from the start, the suspensorium constitutes, with Meckel's cartilage and the hyoid bar, a single cartilaginous element, and the junction between the front and rear of the neurocranium is complete on day 4. By day 8 the quadratomandibular joint has formed and the tectum posterius has appeared. Cartilage reduction first affects the trabecular bars, then, markedly, the visceral arches. By day 18 the braincase floor has almost disappeared.     

The cranial morphology of Greererpeton burkemorani Romer (Amphibia: Temnospondyli)

The skull of Greererpeton burkemorani Romer, a temnospondyl amphibian from the Upper Mississippian at Greer, West Virginia is described. A detailed account of the stapes of a Mississippian amphibian is given for the first time and its function is discussed. It is suggested that the stapes formed the principal element of support for the back of the braincase and resisted potential dislocation of the otico-occipital region from the skull roof during contraction of the hypaxial musculature.
Greererpeton is included in the Colosteidae and an amended diagnosis of the family is given. Erpetosaurus differs from Colosteus, Greererpeton and Pholidogaster in the pattern of bones in the skull roof and palate, the dentition and the otic region and, consequently, it is removed from the Colosteidae. The Temnospondyli are considered to be a monophyletic group characterized by the development of a connection between the dorsal portion of the occipital arch, the exoccipital bones, and the skull roof. The loxommatids are removed from the Temnospondyli as they retain the plesiomorphic condition of braincase attachment which relies exclusively on derivatives of the auditory capsules.
On the basis of similarities in the structure of the braincase, palate and manus it is suggested that microsaurs are the collateral descendants (sister group) of temnospondyls. This relationship may account for the large number of similarities in the three living groups of Amphibia: Anura are generally believed to have descended from temnospondyls, while the Urodela and Apoda are often considered to have descended from microsaurs. These systematic conclusions endorse the recent suggestions that neither the Lepospondyli nor the Labyrinthodontia are natural groups, and both terms should be abandoned.     

Early development of the head skeleton in Brycon moorei(Pisces, Ostariophysi, Characidae)

At hatching (15 h post fertilization), Brycon moorei possesses no skeletal structure. Thereafter, development is very rapid. The first oral teeth appear no later than 3 h post‐hatching, but they remain covered with epithelium until c . 45 h. At 7 h, the trabecular bars and part of the cartilaginous visceral arches are visible and at 15 h, the dentaries and premaxillaries are present. At 25 h, i.e . the onset of piscivory and cannibalism (the yolk sac is only fully resorbed after 36 h), the oral teeth are fully developed, the first pharyngeal teeth are formed, and some head movements already appear synchronized, but the mouth cavity is not completely isolated from the neurocranium by bony structures. Thereafter, no new buccal or pharyngeal bony structure is visible until 45 h, when the maxilla and opercula appear, along with a new type of cannibalistic behaviour. Cartilage resorptions also start at 45 h, but with no concomitant replacement by formation of calcified structures. Later, development gradually becomes similar to that of many previously studied teleosts. The developmental pattern of B. moorei is thus extremely rapid in comparison with other teleosts, i.e . it prioritizes feeding structures that permit the expression of piscivory at a very early age. The uniqueness of this pattern is discussed in relation to ecological constraints on early feeding and fast growth.     

Cranial ontogeny of otariid seals

Over 2000 otariid skulls were measured for a morphometric study of cranial ontogeny in fur seals and sea lions. Few interspecific differences in cranial ontogeny were observed in the Otariidae, with only minor differences in rates of growth. Sexual dimorphism was significant in all otariids but was more apparent in the larger species. Female otariids of each species showed monophasic development in all characteristics, whereas males expressed monophasic growth for some characters and biphasic growth for others. Biphasic development in skulls of male otariids occurred well after physical maturity had been reached, usually at a suture index of 27. The rate of development varied between skull characters; components relating to the nervous system completed growth well before the rest of the skull, whereas those related to feeding, respiration and vocalisation developed in synchrony with the overall growth of total skull length. Sutures of the calvaria, or braincase, were the first to show partial closure and those uniting the facial bones were usually the last to fuse. As with the calvaria, the orbits and otic capsules developed quickly, suggesting a need for good hearing and vision early in independent postnatal life. Development of the rostral and palatal regions required significantly longer to complete growth.     

Skull deformations in craniosynostosis and endocrine disorders: Morphological and tomographic analysis of the skull from the crypt of the Silesian Piasts in Brzeg (16th–17th century), Poland

Results

of morphological and tomographic (CT) studies of the skull that was found in the crypt of the Silesian Piasts in the St. Jadwiga church in Brzeg (Silesia, Poland) are presented and discussed here. The established date of burial of probably a 20–30 years old male was 16th–17th century. The analyzed skull showed premature obliteration of the major skull sutures. It resulted in the braincase deformation, similar to the forms found in oxycephaly and microcephaly. Tomographic analysis revealed gross pathology. Signs of increased intracranial pressure, basilar invagination and hypoplasia of the occipital bone were observed. Those results suggested the occurrence of the very rare Arnold-Chiari syndrome. Lesions found in the sella turcica indicated the development of pituitary macroadenoma, which resulted in the occurrence of discreet features of acromegaly in the facial bones. The studied skull was characterized by a significantly smaller size of the neurocranium (horizontal circumference 471 mm, cranial capacity ∼1080 ml) and strongly expressed brachycephaly (cranial index = 86.3), while its height remained within the range for non-deformed skulls. A narrow face, high eye-sockets and prognathism were also observed. Signs of alveolar process hypertrophy with rotation and displacement of the teeth were noted. The skull showed significant morphological differences compared to both normal and other pathological skulls such as those with pituitary gigantism, scaphocephaly and microcephaly.     

Anatomical study of the skull of amphisbaenian Diplometopon zarudnyi (Squamata,Amphisbaenia)

This study investigates the amphisbaenian species skull which includes cranium, lower jaw and hyoid apparatus. The medial dorsal bones comprise the premaxilla, nasal, frontal and parietal. The premaxilla carries a large medial tooth and two lateral ones. The nasals are paired bones and separated by longitudinal suture. Bones of circumorbital series are frontal, orbitosphenoid and maxilla. The occipital ring consists of basioccipital, supraoccipital and exooccipital. Supraoccipital and basioccipital are single bones while the exo-occipitals are paired. The bones of the palate comprise premaxilla, maxilla, septomaxilla, palatine, pterygoid, ectopterygoid, basisphenoid, parasphenoid, orbitosphenoid and laterosphenoid. Prevomer and pterygoid teeth are absent. Palatine represent by two separate bones. The temporal bones are clearly visible. The lower jaw consists of the dentary, articular, coronoid, supra-angular, angular and splenial. The hyoid apparatus is represented by a Y-shaped structure. The mandible is long and is suspended from the braincase via relatively short quadrate. There is an extensive contact between the long angular and the large triangular coronoid. Thus inter-mandibular joint is bridged completely by the angular and consequently, the lower jaws are relatively rigid and kinetic. The maxillae are suspended from the braincase largely by ligaments and muscles rather than through bony articulation. In conclusion, the skull shape affects feeding strategy in Diplometopon zarudnyi. The prey is ingested and transported via a rapid maxillary raking mechanism.     

Early development of the chondrocranium in Salmo letnica(Karaman, 1924)(Teleostei: Salmonidae)

The ontogenetic development of the chondrocranium of Ohrid trout Salmo letnica was studied from hatching until 92 days post‐hatching (dph). Most of the samples were in toto trypsin cleared and stained, some specimens were used for serial histological sectioning. The serial histological sections of fish specimens at the age of 92 dph were used for a graphical reconstruction of the cartilaginous neurocranium. A chronological evaluation of the formation of the cartilaginous skull in the early development of S. letnica was performed. In order to investigate to what degree the ontogeny of the Ohrid trout is unique, the results were compared with data of the development of other salmonids, as well as some non‐salmonid teleosts. The development of the cartilaginous structures of the Ohrid trout was found to be similar to that of other salmonids. Most of the cartilage structures of the neurocranium and the viscerocranium are present at the moment of hatching of this species. A fully developed chondrocranium was observed at the age of 92 dph, when the first signs of cartilage resorption could also be observed.     

Cranial osteogenesis in Monodelphis domestica (Didelphidae) and Macropus eugenii (Macropodidae)

The pattern of onset and general rate of cranial ossification are compared in two marsupials, Monodelphis domestica (Didelphidae) and Macropus eugenii (Macropodidae). In both species a similar suite of bones is present at birth, specifically those surrounding the oral cavity and the exoccipital, and in both postnatal events follow a similar course. The facial skeleton matures more rapidly than the neurocranium, which is characterized by an extended period of ossification. Most dermal bones begin ossification before most endochondral bones. Endochondral bones of the neurocranium are particularly extended in both the period of onset of ossification and the rate of ossification. These data confirm suggestions that morphology at birth is conservative in marsupials and we hypothesize that the pattern of cranial osteogenesis is related to two distinct demands. Bones that are accelerated in marsupials are correlated with a number of functional adaptations including head movements during migration, attachment to the teat, and suckling. However, the very slow osteogenesis of the neurocranium is probably correlated with the very extended period of neurogenesis. Marsupials appear to be derived relative to both monotreme and placental mammals in the precocious ossification of the bones surrounding the oral cavity, but share with monotremes an extended period of neurocranial osteogenesis. © 1993 Wiley-Liss, Inc.     

Cranial anatomy of the extinct amphisbaenian Rhineura hatcherii (Squamata, Amphisbaenia) based on high-resolution X-ray computed tomography

The fossilized skull of a small extinct amphisbaenian referable to Rhineura hatcherii Baur is described from high-resolution X-ray computed tomographic (HRXCT) imagery of a well-preserved mature specimen from the Brule Formation of Badlands National Park, South Dakota. Marked density contrast between bones and surrounding matrix and at bone-to-bone sutures enabled the digital disarticulation of individual skull elements. These novel visualizations provide insight into the otherwise inaccessible three-dimensionally complex structure of the bones of the skull and their relationships to one another, and to the internal cavities and passageways that they enclose. This study corrects several previous misidentifications of elements in the rhineurid skull and sheds light on skull construction generally in "shovel-headed" amphisbaenians. The orbitosphenoids in R. hatcherii are paired and entirely enclosed within the braincase by the frontals; this is in contrast to the condition in many extant amphisbaenians, in which a large azygous orbitosphenoid occupies a topologically distinct area of the skull, closing the anterolateral braincase wall. Rhineura hatcherii retains a vestigial jugal and a partially fused squamosal, both of which are absent in many extant species. Sculpturing on the snout of R. hatcherii represents perforating canals conveying sensory innervation; thus, the face of R. hatcherii receives cutaneous innervation to an unprecedented degree. The HRXCT data (available at www.digimorph.org) corroborate and extend previous hypotheses that the mechanical organization of the head in Rhineura is organized to a large degree around its burrowing lifestyle.     

Development of the cartilaginous skull in solea solea: trends in pleuronectiforms

The postembryonic development of the cartilaginous cephalic skeleton in Solea solea (Linné) was observed from hatching to the juvenile stage or postmetamorphic larva. Fry were trypsin-cleared and alcian-blue-stained. At hatching, Solea solea displays no cartilaginous structure. On day 3, the fry exhibit trabecular bars, Meckel's cartilages, palatoquadrates, hyosymplectics, hyoid bars, the first two ceratobranchials, and basibranchials 1–3. On day 4, ceratobranchials 3 and 4 form. On day 12, short taeniae marginales are formed on the well-developed otic capsules and the branchial basket is nearly complete. On day 14, a first cartilage reduction begins as the posterior part of the trabecula communis separates from the otic capsules. On day 18, metamorphosis begins. The tectum posterius completes posteriorly the braincase dome. On day 20, the left eye moves to the median crest of the head and the right eye shifts downwards slightly. Consequently, displacement of the right eye induces a deformation extending to the right pterygoid process. On day 23, both eyes are located on the same side. On day 29, cartilage reduction affects the hyosymplectics; it continues with the first two ceratobranchials on day 32 and with the otic capsules on day 40. On day 50, cartilage reduction has reached the lamina precerebralis. Only a few small lumps remain at the level of the neurocranium and the splanchnocranium.     

Covariation of brain and skull shapes as a model to understand the role of crosstalk in development and evolution

Covariation among discrete phenotypes can arise due to selection for shared functions, and/or shared genetic and developmental underpinnings. The consequences of such phenotypic integration are far-reaching and can act to either facilitate or limit morphological variation. The vertebrate brain is known to act as an “organizer” of craniofacial development, secreting morphogens that can affect the shape of the growing neurocranium, consistent with roles for pleiotropy in brain–neurocranium covariation. Here, we test this hypothesis in cichlid fishes by first examining the degree of shape integration between the brain and the neurocranium using three-dimensional geometric morphometrics in an F₅ hybrid population, and then genetically mapping trait covariation using quantitative trait loci (QTL) analysis. We observe shape associations between the brain and the neurocranium, a pattern that holds even when we assess associations between the brain and constituent parts of the neurocranium: the rostrum and braincase. We also recover robust genetic signals for both hard- and soft-tissue traits and identify a genomic region where QTL for the brain and braincase overlap, implicating a role for pleiotropy in patterning trait covariation. Fine mapping of the overlapping genomic region identifies a candidate gene, notch1a, which is known to be involved in patterning skeletal and neural tissues during development. Taken together, these data offer a genetic hypothesis for brain–neurocranium covariation, as well as a potential mechanism by which behavioral shifts may simultaneously drive rapid change in neuroanatomy and craniofacial morphology.     

The influence of experimental deformation on craniofacial development in rats

Cranial deformation was produced experimentally in rats 8 to 40 days old for the purpose of studying the rotation of the craniofacial bones and the modification of the growth rates of the functional cranial components. One hundred and twenty four skulls (65 males and 59 females) were employed, classified as: deformed , deformed-hydrocephalic, sham-operated and controls. A midsagittal diagram was drawn for each skull and the angle subtended by each bone with respect to the vestibular plane was measured. Growth indices were worked out for both the neural skull and the facial skull. Deformation altered the rotation of the parietal, interparietal and basisphenoidal bones and restricted the rotation of the fronto-ethmo-facial complex. Alteration of the longitudinal growth rates of the dorsal and basilar components of the neurocranium and the splanchnocranium produced the persistence of the klinorynchal state.